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Primary Production in Coastal Lagoons - A COASTAL ZONES AND ESTUARIES – Primary Production in Coastal Lagoons - A. Vázquez-Botello, F. Contreras-Espinosa, G. De La Lanza-Espino and S. Villanueva F. PRIMARY PRODUCTION IN COASTAL LAGOONS A. Vázquez-Botello Instituto de Ciencias del Mar, UNAM, México F. Contreras-Espinosa Universidad Autónoma Metropolitana-Iztapalapa, México G. De La Lanza-Espino Instituto de Biología, UNAM. México S. Villanueva F. Instituto de Ciencias del Mar, UNAM. México. Keywords: primary production, primary productivity, coastal lagoons, tropical estuaries, submerged vegetation, mangroves, marshes, estuarine phytoplankton, seagrasses, coastal wetlands, chlorophyll “a”. Contents 1. Introduction 2. Measurement of primary production 3. Temporal and spatial variations 4. Physical setting and primary producers 5. Primary production in Mexican coastal lagoons 5.1. Role of macrovegetation in littoral production 5.1.1 Saltmarshes 5.1.2 Submerged Vegetation 5.1.3 Mangrove 6. Conclusions Acknowledgements Glossary Bibliography Biographical Sketches 1. Introduction UNESCO – EOLSS The expressions primary production and productivity have been used indiscriminately in aquatic environments,SAMPLE even to the extent CHAPTERSof using the same measuring units, with some minor differences. According to Wetzel (1975) production is the weight of new organic matter formed in a period of time per volume or area, plus losses due to respiration, excretion, secretion, damage, death, or grazing. It is also the increase in biomass in a given time. Primary productivity is the potential rate of biomass addition per time per area; this term implies an average instantaneous rate per hour, per day, or per year. These two concepts have been confused with biomass, which differs basically in the units: biomass is not a rate, but it is the weight of plant and animal matter in a given area; hence, it can be expressed in g/m2, g/m3, or volume displaced per liter. ©Encyclopedia of Life Support Systems (EOLSS) COASTAL ZONES AND ESTUARIES – Primary Production in Coastal Lagoons - A. Vázquez-Botello, F. Contreras-Espinosa, G. De La Lanza-Espino and S. Villanueva F. The primary producers, photosynthesizers or autotrophic organisms of aquatic environments encompass microscopic organisms (phytoplankton), macroalgae, and phanerogams (from very small free-floating plants to large trees, such as mangroves. They all participate in inorganic carbon assimilation to produce organic carbon at diverse rates. Although, published data on primary production and/or productivity in coastal lagoons has increased in the last years, their interpretation still poses a problem since many studies have focused mainly on the description of the most relevant producing populations. Primary production in coastal lagoons can be dominated by phytoplankton, benthic micro- and/or macro-algae, macrophytes, and, in special cases, by a combination of all of them. The afore-mentioned emphasizes the need for new and diverse sampling strategies and analytical methods, both spatial and temporal, as well as a balanced interpretation, to cover the existing information gaps. Being the ground stock of the food chains, primary production must be assessed precisely from both the ecological and economic (e.g. fisheries) standpoints, not only because of the knowledge per se but also for the management and preservation of aquatic environments. 2. Measurement of primary production Determination of phytoplanktonic primary production in tropical coastal lagoons has become controversial, with some ecological principles based on data from temperate latitudes (Russell-Hunter, 1970; Odum, 1972; Margalef, 1974; Goldman, 1974; Vollenweider, 1974). Numerous authors have dealt with the measurement and expression of primary production since the classical works of Rhyther in the 1950s. One of the greatest challenges has been to establish a worldwide reliable and compatible methodology. For the study of phytoplanktonic photosynthesis there are essentially four methods for micro- and macro-vegetation with some variations, based on: 1. Production of oxygen in clear and dark bottles (Gaarder and Gran, 1927). The methodological problem with the dark bottle due to excessive light has been solved by using a photosynthetic inhibitor (DCMU) (Legendre et al., 1983). This determinationUNESCO includes also the diurna– l EOLSSoxygen curve of Odum (1956) based on the production of oxygen or its consumption by morning photosynthesis or nocturnal SAMPLErespiration and their equilibrium CHAPTERS in open or in situ environments. This evaluation is performed by sampling the water every two or three hours to determine dissolved oxygen in a 24-hour period (Leith, 1975; Whittaker, 1975). 2. Incorporation rate of C14 radioactive carbon (Steeman Nielsen, 1952). An under- estimation implicit in the C14 methodology is caused by the excretion products that are usually not assessed and which can represent from 0 to 99% of the production, according to the stress caused by the environmental conditions on the organisms (de la Lanza-Espino and Lozano, 1999). 3. Rate of assimilated CO2 evaluated by alkalinity and pH changes (Beyers and Odum, 1960). ©Encyclopedia of Life Support Systems (EOLSS) COASTAL ZONES AND ESTUARIES – Primary Production in Coastal Lagoons - A. Vázquez-Botello, F. Contreras-Espinosa, G. De La Lanza-Espino and S. Villanueva F. 4. Rate of increases in particulate matter per time (Rhyther and Menzel, 1965). Determination of chlorophyll “a” associated to appropriate light, although not implying time, is an inexpensive index of the productivity of an environment. A range of 0.9 to 3.0 mg C/mg of chlorophyll “a”, is considered a typical value; this quotient varies daily, with a maximum in the morning, a decrease at noon, and a minimum at night (Goldman et al., 1963; Mc Allister, 1963). However, Hall and Moll (1975), among other authors, consider that its determination only enables speculation about photosynthesis, and that it results not only from spatial and temporal variations but also depends on the efficiency of the pigment, since it could be inactive and not proportional to the size and age of the cells. It has been stated that results with chlorophyll “a” are not better than equations used to predict photosynthesis. However, the other techniques, except for the daily oxygen curve, also have disadvantages as a result of isolating the organisms from their natural environment. Most phytoplankton cells have other pigments, in addition to chlorophyll “a”; these are known as accessories and they serve as energy transporters (Golterman, 1975), protecting chlorophyll “a” from light (Salisbury and Ross, 1969). The ratio of chlorophyll “a” to carotenoids depends basically on the physiological conditions of the cell (Yentsch, 1965). Margalef (1981) relates primary productivity with the structure of the community through pigment proportions to obtain an index of Species Diversity (D430/D665), which not only reflects changes in the composition of the species in the community but also, the physiological state of the population. From a technical point of view, the concept of Productivity Index (PI) was introduced in the 1960s. This index considers growth kinetics of unicellular organisms (marine phytoplankton), which has been poorly appreciated (Strickland, 1966). This is mainly due to the fact that the primary net production of the phytoplankton provides little information on the intensity of the production or vitality of a population: large biomasses can photosynthesize with lower rates and small biomasses can have high production rates. Therefore a better understanding of biomass diversity and net production of endemic populations is necessary. Hence, in order to be precise, evolution of the net photosynthetic rate per biomass unit at an optimal light level has to be known (0.1 to 0.15 Langley/min). Therefore, PI is expressed as: PI = Production rate per biomass (at one unit of light intensity) UNESCO Amount of biomass – EOLSS = dp x 1 = K SAMPLEat one unit of light intensity CHAPTERS dt where: p = biomass t = time K = constant ©Encyclopedia of Life Support Systems (EOLSS) COASTAL ZONES AND ESTUARIES – Primary Production in Coastal Lagoons - A. Vázquez-Botello, F. Contreras-Espinosa, G. De La Lanza-Espino and S. Villanueva F. This allows comparing PI values at relevant luminic intensities and the amount of carbon associated with the cells. It must be emphasized that evaluations must be made according to the time of division and growth of the organisms. Unfortunately, each of these methods yields different results, including measurement units (gC/m2/h, gC/m2/d, gC/m2/year, or the same but using m3), hindering comparisons (Hall and Moll, 1975; Peterson, 1980; de la Lanza-Espino et al., 1991). The first two methods are the most currently used to measure photosynthesis, and although the bottle method has been valued, some experiments comparing C14 and oxygen evolution methods have provided similar results (Williams et al., 1979, 1983). It is possible to state that in estuarine-lagoon systems, the photosynthetic process in the water column is complex and highly specific in terms of localization. Variations in the same day and site are extreme, which shed doubts on any type of generalization even for the same area (Contreras, 1995). In contrast, the chlorophyll “a” content has been proven to be more constant in any situation, making it a more reliable factor. The relation between chlorophyll “a” and primary productivity, for some authors the "Assimilation Index" (Beerman
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